Transcript AnimalDiversity3

Most animal phyla
originated in a brief span
of geological time
• The fossil record and molecular studies concur
that the diversification that produced most animal
phyla occurred rapidly on the vast scale of
geologic time.
• This lasted about 40 million years (about 565 to
525 million years ago) during the late
Precambrian and early Cambrian (which began
about 543 million years ago).
• The strongest evidence for the initial
appearance of multicellular animals is found
in the the last period of the Precambrian
era, the Ediacaran period.
– Fossils from the Ediacara Hills of Australia (565 to
543 million years ago) and other sites around the
world consist primarily of cnidarians, but softbodied mollusks were also present, and
numerous fossilized burrows and tracks indicate
the presence of worms.
– Recently, fossilized animal embryos in China from
570 million years ago and what could be
fossilized burrows from rocks 1.1 billion years
ago have been reported.
• Data from molecular systematics suggest an
animal origin about a billion years ago.
• Nearly all the major animal body
plans appear in Cambrian rocks from
543 to 525 million years ago.
• During this relatively short time, a
burst of animal origins, the
Cambrian explosion, left a rich
fossil assemblage.
– It includes the first animals with hard,
mineralized skeletons
• On the scale of geologic time, animals
diversified so rapidly that it is difficult
from the fossil record to sort out the
sequence of branching in animal
phylogeny.
– Because of this, systematists depend
largely on clues from comparative
anatomy, embryology, developmental
genetics, and molecular systematics of
extant species.
• Some systematists studying animal
phylogeny interpret the molecular
data as supporting three Cambrian
explosions, not just one.
• For the three main branches of
bilateral animals - Lophotrochozoa,
Ecdysozoa, and Deuterostomia - the
relationships among phyla within each
are difficult to resolve, but the
differences between these three clades
are clear, based on their nucleic acid
sequences.
– This suggests that these three clades
branched apart very early, probably in the
Precambrian.
• Zoologists recognize about 35 phyla of
animals.
• For the past century, there was broad
consensus among systematists for the major
branches of the animal phylogenetic tree.
– This was based mainly on anatomical features in
adults and certain details of embryonic
development.
• However, the molecular systematics of the
past decade is challenging some of these
long-held ideas about the phylogenetic
relationships among the animal phyla.
The remodeling of phylogenetic trees
illustrates the process of scientific
inquiry
• It can be frustrating that the phylogenetic trees in
textbooks cannot be memorized as infallible truths.
• On the other hand, the current revolution in
systematics is a healthy reminder that science is
both a process of inquiry and dynamic.
– Emerging technologies such as molecular biology
and fresh approaches such as cladistics produce
new data or stimulate reconsideration of old data.
• New hypotheses or refinements of old
ones represent the latest versions of
what we understand about nature
based on the best available evidence.
• Evidence is the key word because
even our most cherished ideas in
science are probationary.
– Science is partly distinguished from other
ways of knowing because its ideas can
be falsified through testing with
experiments and observations.
– The more testing that a hypothesis
withstands, the more credible it
becomes.
• A comparison of the traditional
phylogenetic tree of animals with the
remodeled tree based on molecular
biology shows agreement on some
issues and disagreement on others.
• Though the new data from molecular
systematics are compelling, the
traditional view of animal phylogeny
still offers some important
advantages for helping us understand
the diversity of animal body plans.
The traditional phylogenetic tree
of animals is based mainly on
grades in body “plans”
• The traditional view of relationships
among animal phyla are based mainly
on key characteristics of body plans
and embryonic development.
• Each major branch represents a grade,
which is defined by certain body-plan
features shared by the animals
belonging to that branch.
• The basic organization of germ layers,
concentric layers of embryonic tissue that
form various tissues and organs, differs
between radiata and bilateria.
• The radiata are said to be diploblastic
because they have two germ layers.
– The ectoderm, covering the surface of the
embryo, give rise to the outer covering and, in
some phyla, the central nervous system.
– The endoderm, the innermost layer, lines the
developing digestive tube, or archenteron, and
gives rise to the lining of the digestive tract and
the organs derived from it, such as the liver and
lungs of vertebrates.
• The bilateria are triploblastic.
– The third germ layer, the mesoderm lies
between the endoderm and ectoderm.
– The mesoderm develops into the muscles
and most other organs between the
digestive tube and the outer covering of
the animal.
(3) The Bilateria can be divided by the
presence or absence of a body cavity (a
fluid-filled space separating the digestive
tract from the outer body wall) and by the
structure the body cavity.
• Acoelomates (the phylum
Platyhelminthes) have a solid body and
lack a body cavity.
• In some organisms, there is a body
cavity, but it is not completely lined
by mesoderm.
– This is termed a pseudocoelom.
– These pseudocoelomates include the
rotifers (phylum Rotifera) and the
roundworms (phylum Nematoda).
• Coelomates are organisms with a
true coelom, a fluid-filled body cavity
completely lined by mesoderm.
– The inner and outer layers of tissue that
surround the cavity connect dorsally and
ventrally to form mesenteries, which
suspend the internal organs.
• A body cavity has many functions.
– Its fluid cushions the internal organs,
helping to prevent internal injury.
– The noncompressible fluid of the body
cavity can function as a hydrostatic
skeleton against which muscles can
work.
– The presence of the cavity enables the
internal organs to grow and move
independently of the outer body wall.
(4) The coelomate phyla are divided
into two grades based on differences
in their development.
– The mollusks, annelids, arthropods, and
several other phyla belong to the
protostomes, while echinoderms,
chordates, and some other phyla belong
to the deuterostomes.
– These differences center on cleavage
pattern, coelom formation, and
blastopore fate.
• Protostomes generally undergo spiral
cleavage, in which planes of cell division
are diagonal to the vertical axis of the
embryo.
– Most protostomes also show determinate
cleavage where the fate of each embryonic cell
is determined early in development.
• The zygotes of deuterostomes undergo
radial cleavage in which the cleavage
planes are parallel or perpendicular to the
vertical egg axis.
– Deuterostomes show indeterminate cleavage
whereby each cell in the early embryo retains
the capacity to develop into a complete embryo.
• Coelom formation begins in the
gastrula stage.
– As the archenteron forms in a
protostome, solid masses of mesoderm
split to form the coelomic cavities, called
schizocoelous development.
– In deuterostomes, mesoderm buds off
from the wall of the archenteron and
hollows to become the coelomic cavities,
called enterocoelous development.
• The third difference centers on the
fate of the blastopore, the opening
of the archenteron.
– In many protosomes, the blastopore
develops into the mouth and a second
opening at the opposite end of the
gastrula develops into the anus.
– In deuterostomes, the blastopore usually
develops into the anus and the mouth is
derived from the secondary opening.
• Molecular systematics has added a
new set of shared-derived characters
in the form of unique monomer
sequences within certain genes and
their products.
– These molecular data can be used to
identify the clusters of monophyletic taxa
that make up clades.
• In some cases, the clades determined
from molecular data reinforce the
traditional animal tree based on
comparative anatomy and
development, but in other cases, a
very different pattern emerges.
• This
phylogenetic
tree is based
on nucleotide
sequences
from the
small subunit
ribosomal
RNA.
• At key places, these two views of
animal phylogeny are alike.
– First, both analyses support the
traditional hypotheses of the ParazoaEumetazoa and Radiata-Bilateria
dichotomies.
– Second, the molecular analysis reinforces
the hypothesis that the deuterostomes
(echinoderms and chordates) form a
clade.
• However, the traditional and
molecular-based phylogenetic trees
clash, especially on the protostome
branch.
– The molecular evidence supports two
protostome clades: Lophotrochozoa,
which includes annelids (segmented
worms) and mollusks (including clams
and snails), and Ecdysozoa, which
includes the arthropods.
• Traditional analyses have produced
two competing hypotheses for the
relationships among annelids,
mollusks, and arthropods.
– Some zoologists favored an annelidarthropod lineage, in part because both
have segmented bodies.
– Other zoologists argued that certain
features favored an annelid-mollusk
lineage, especially because they share a
similar larval stage, the trochophore
larva.
• This hypothesis is supported by the molecular
data.
• Traditionally, the acoleomate phylum
Platyhelminthes (flatworms) branches from
the tree before the formation of body
cavities.
– The molecular data place the flatworms within
the lophotrochozoan clade.
– If this is correct, then flatworms are not
primitive “pre-coelomates” but are protostomes
that have lost the coelom during their evolution.
• The molecular-based phylogeny splits the
pseudoceolomates with the phylum Rotifera
(rotifers) clustered with the
lophotrochozoan phyla and the phylum
Nematoda (nematodes) with the
ecdysozoans.
• In the traditional tree, the assignment
of the three lophophorate phyla is
problematic.
– These animals have a lophophore, a
horseshoe-shaped crown of ciliated tentacles
used for feeding.
– The lophophorate phyla share
some characteristics with
protostomes and other features
with deuterostomes.
– The molecular data place the
lophophorate phyla among the
phyla with the trochophore
larvae, hence the name
lophotrochozoans.
• The name Ecdysozoa (nematodes,
arthropods, and other phyla) refers to
animals that secrete external
skeletons (exoskeleton).
– As the animal grows, it molts the old
exoskeleton and secretes a new, larger one, a
process called ecdysis.
– While named for this
process, the clade is
actually defined mainly
by molecular evidence.
– In summary, the molecular evidence recognizes
two distinct clades within the protostomes and
distributes the acoelomates, pseudocoelomates,
and lophophorate phyla among these two clades.
• Our survey of animal phyla is based on the
newer molecular phylogeny, but there are
two caveats.
• First, the concept of body-plan grades still
is a very useful way to think about the
diversity of animal forms that have
evolved.
• Second, the molecular phylogeny is a
hypothesis about the history of life, and is
thus tentative.
– This phylogeny is based on just a few genes mainly the small subunit ribosomal RNA (SSUrRNA).
– Ideally, future research, including fossil
evidence and traditional approaches, will
eventually square the molecular data with data
from these other approaches.